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Title Feeding Characteristics of a Japanese Pitviper, Ovophis Feeding Characteristics of a Japanese Pitviper, Ovophis Title okinavensis, on Okinawa Island: Seasonally Biased but Ontogenetically Stable Exploitation on Small Frogs Author(s) MORI, Akira; TODA, Mamoru Citation Current Herpetology (2011), 30(1): 41-52 Issue Date 2011-07 URL http://hdl.handle.net/2433/197271 Right © 2011 The Herpetological Society of Japan Type Journal Article Textversion publisher Kyoto University Current Herpetology 30(1): 41–52, June 2011 © 2011 by The Herpetological Society of Japan Feeding Characteristics of a Japanese Pitviper, Ovophis okinavensis, on Okinawa Island: Seasonally Biased but Ontogenetically Stable Exploitation on Small Frogs 1 1,2 AKIRA MORI * AND MAMORU TODA 1 Department of Zoology, Graduate School of Science, Kyoto University, Sakyo, Kyoto 606–8502, JAPAN 2 Present address: Tropical Biosphere Research Center, University of the Ryukyus, Nishihara, Okinawa 903–0213, JAPAN Abstract: Ovophis okinavensis is an endemic pitviper of the subtropical region of Japan. Based on information from various localities, the species has been considered a dietary generalist that exploits various terrestrial vertebrates. We analyzed stomach contents of O. okinavensis in a northern mountain area on Okinawa Island, based on data of a 10-years-study, to examine its trophic features at the population level. We recovered a total of 461 prey items, which were comprised of six, three, five, and three species of frogs, reptiles, birds, and mammals, respectively. Approximately 93% of dietary items consisted of frogs, the majority of which belonged to two explosively breeding species aggregating to restricted spots along the stream only during their reproductive season (winter). On the other hand, from April to November, the snake mainly depended on vertebrates other than frogs. Neither an ontogenetic dietary shift nor intersexual dietary divergence was evident although males tended to exploit one of the two explosively breeding frog species more frequently than females. There were weak positive correlations between snake size and prey size, but large snakes did not drop small prey items from their diet. Average body mass of each prey item was only 5.2% of snake body mass. Characteristics of trophic features of O. okinavensis of this population, such as extremely high dependency on frogs in winter, the absence of ontogenetic dietary shift, and utilization of small sized prey, are discussed in comparison with other viperids. Key words: Crotalinae; Prey size; Ontogenetic dietary shift; Ovophis okinavensis; Sexual difference INTRODUCTION exclusive of Australasia and Antarctica (Pough et al., 2004). Since those early works by a few Viperidae is one of the much diversified authors, such as Saint Girons (1952), Fitch snake families widely radiating over the world (1960), and Klauber (1972), the trophic ecol- ogy and feeding characteristics of viperid * Corresponding author. Tel: +81–75–753–4075; snakes have been relatively well studied chiefly Fax: +81–75–753–4075; on the basis of data from European and North E-mail address: [email protected] American species. With respect to members 42 Current Herpetol. 30(1) 2011 of the family from other regions, most compa- okinavensis, such as sexual differences, rable studies started much more recently ontogenetic shift, seasonal change, and effects (South America: Valdujo et al., 2002; Nogueira of snake’s body size on the prey properties. et al., 2003; Monteiro et al., 2006, Central Here, we investigate the feeding habit of O. Africa: Luiselli and Akani, 2003; Ineich et al., okinavensis by analyzing sexual, ontogenetic, 2006, Southeast Asia: Daltry et al., 1998; and seasonal divergences in prey taxa and size, Creer et al., 2002; Mori et al., 2002; Shine and based on a long-term study conducted in the Sun, 2003; Lin and Tu, 2008), but these studies northern mountain area on Okinawa Island. also have revealed their diverse feeding habits We then discuss the trophic features of this as ranging from frog specialists (Ineich et al., pitviper by comparing its general feeding 2006) to vertebrate generalists (Creer et al., habits with those of other viperid snakes 2002; Valdujo et al., 2002; Monteiro et al., previously reported. 2006). These studies, as well as several other studies on some North American species, have MATERIALS AND METHODS also highlighted a possible common tendency of ontogenetic dietary shift, typically from The diet of O. okinavensis was examined on ectothermic to endothermic prey (Klauber, the basis of the stomach contents of individual 1972; Campbell and Lamar, 1989; Daltry et snakes captured from December 1996 to al., 1998; Shine and Wall, 2007; Lin and Tu, February 2006 in a limited area of the north- 2008), along with improvement of ability to ern part of Okinawa Island, Ryukyu Archipel- ingest larger prey through structural modifica- ago, Japan. The main study area (ca. 25 ha) tions of trophic apparatus, an evolutionary encompasses the upper streams of Zatsun innovation characteristic to viperid snakes River and surrounding hilly environments of (Pough and Groves, 1983; Greene, 1992; but Yambaru. Vegetation was dominated by see Cundall and Deufel, 2006). Nonetheless, a primary and well-recovered secondary forests great majority of the Asian and African consisting of humid-subtropical broad-leaved viperids remains to be studied (Ineich et al., evergreen trees such as Castanopsis sieboldii. 2006). Furthermore, dietary information on a Meteorological data of the study region is given Asian or African species, even when shown in Mori et al. (2002). Field study was available, mostly derives from museum speci- made throughout the year but with particular mens originated from various localities: very intensities from December to March for the few population-based studies have been con- purpose of investigating the relationship of a ducted despite their obvious importance for winter activity of the snake with the breeding our better understanding of feeding traits in an activity of two species of frogs, Rana sp. evolutionary framework (Thompson, 2005). (formerly erroneously referred to as R. Ovophis okinavensis is a small, subtropical okinavana, see Matsui [2007]; hereafter pitviper distributed on islands in the central referred to as Rana sp. A) and R. narina, both part of the Ryukyu Archipelago, Japan. Cur- breeding in the stream (Mori et al., 2002, rently available dietary records of wild O. 2009). Basically, we haphazardly walked at okinavensis cover all terrestrial vertebrate night along streams and trails, which run classes (Nakachi, 1991; Mori et al., 2002; Toda though the study area irregularly, searching for et al., 2003; Kadota, 2006, 2011; see Mori and snakes on the surface. Whenever we found Moriguchi, 1988 for other references), imply- individual snakes, we collected them except for ing that this species is a generalist feeder. On the cases specified below. We also searched the other hand, despite the abundance of between rocks, insides of crevices of ledges, dietary records, few studies have examined and in burrows on banks. patterns and causes of variations in feeding In addition to the above surveys, we also characteristics within a single population of O. used a road cruising technique to find snakes MORI & TODA—FEEDING CHARACTERISTICS OF PITVIPER 43 on the road to collect dietary data from differ- scale clipping and by painting numbers on ent habitats. The road is an approximately dorsal surfaces of head and posterior body for 7 km segment of the routes Benoki, Ie, and permanent and temporal visual identifications, Chinufuku, which partially surrounds the respectively. In addition, from August 1998, main study area and crosses the Yambaru small (11.5×2.2 mm) passive integrated tran- forests. These routes run through various sponder (PIT) tags were injected under the habitats, such as forests along hill ridges, skin for individual identification. Each snake slopes, and streams, and a forest park with was released at the site of capture basically more open habitats. The straight distance within 24 hours after collection. from the main study area to the farthest point When we found snakes that had been of the segment is approximately 3 km. The collected and released within a few weeks width of the paved road varied from 5 to 8 m. before (visually determined by checking the We drove at 20 to 40 km/h, depending on painting), we usually did not collect them or visibility, and collected snakes whenever we did not examine stomach contents so as not to found them on the way to and back from the disturb them too frequently. Prey animals main study area. confirmed by direct observations on predation Snakes were brought to the nearby field were also recorded as food items. station within 12 hours of collection, where In our earlier study, only a few snakes were they were measured for snout-vent length found along the stream by visual census (SVL), body mass (BM), maximum head width between late spring and late fall (Mori et al., (MHW), interocular distance (IOD: distance 2002). We therefore periodically examined between exterior edges of eyes), and upper jaw between March 2004 and February 2005 length (JL: straight distance between the tip of stomach contents of four radio-tracked snakes the snout and the posterior edge of the eighth (two males and two females), to which we supralabial). MHW and JL were measured surgically implanted radio tags (Holohil SB- while the snakes were gently held by the neck 2T, 5 g) intraperitoneally following the proce- region so as to keep the head slightly above dures described by Reinert and Cundall (1982) the substrate and the mouth closed. Although and Nishimura et al. (1995). These snakes JL is not a direct measurement of the upper were initially collected along the stream and jaw, we used this measurement as one of the released at the site of capture four to six days approximate indices of head size. Sex was after surgery. We located the radio-tracked determined by using a probe, by everting hemi- snakes once every two weeks and collected penes, or by examining the external shape of them to examine the presence of the stomach the tail base.
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